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, 548 (7666), 214-218

Genetic Origins of the Minoans and Mycenaeans

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Genetic Origins of the Minoans and Mycenaeans

Iosif Lazaridis et al. Nature.

Abstract

The origins of the Bronze Age Minoan and Mycenaean cultures have puzzled archaeologists for more than a century. We have assembled genome-wide data from 19 ancient individuals, including Minoans from Crete, Mycenaeans from mainland Greece, and their eastern neighbours from southwestern Anatolia. Here we show that Minoans and Mycenaeans were genetically similar, having at least three-quarters of their ancestry from the first Neolithic farmers of western Anatolia and the Aegean, and most of the remainder from ancient populations related to those of the Caucasus and Iran. However, the Mycenaeans differed from Minoans in deriving additional ancestry from an ultimate source related to the hunter-gatherers of eastern Europe and Siberia, introduced via a proximal source related to the inhabitants of either the Eurasian steppe or Armenia. Modern Greeks resemble the Mycenaeans, but with some additional dilution of the Early Neolithic ancestry. Our results support the idea of continuity but not isolation in the history of populations of the Aegean, before and after the time of its earliest civilizations.

Figures

Extended Data Figure 1
Extended Data Figure 1. ADMIXTURE analysis
ADMIXTURE analysis with K=2 to K=17 is shown. 351 ancient and 2,616 present-day individuals were used in this analysis; ancient samples and present-day Greeks are displayed. To avoid visual clutter of labels, individuals in populations with sample size ≤5 are shown with thicker lines.
Extended Data Figure 2
Extended Data Figure 2. Symmetry testing of Aegean Bronze Age populations
The statistic f4(X, Y; Test, Chimp) is shown with ±3 standard errors. Each panel is titled with the pair X, Y. Populations are ordered according to the value of the statistic. Positive values indicate that Test shares more alleles with X than Y and negative values that it shares more with Y than X. (a) ‘northern’ and ‘eastern’ populations share more alleles with Minoans than with Neolithic Greece. (b) ‘northern’ and ‘eastern’ populations share more alleles with Mycenaeans than with Neolithic Greece. (c) Minoans from Lasithi and Moni Odigitria are symmetrically related to diverse populations. (d) Neolithic populations from Anatolia, Europe, Greece, and the Levant share fewer alleles with Mycenaeans than with Minoans.
Extended Data Figure 3
Extended Data Figure 3. Symmetry testing of Anatolian Bronze Age populations
The statistic f4(X, Y; Test, Chimp) is shown with ±3 standard errors. Each panel is titled with the pair X, Y. Populations are ordered according to the value of the statistic. Positive values indicate that Test shares more alleles with X than Y and negative values that it shares more with Y than X. (a) European, Siberian, and Caucasus hunter-gatherers share fewer alleles with Bronze Age Anatolians from Harmanören Göndürle than with a Chalcolithic Anatolian from Barcın. (b) Bronze Age Anatolians differ from Neolithic ones in sharing more alleles with populations of Iran, the Caucasus, and the Steppe than with those of Europe. (c) Bronze Age Anatolians differ from Minoans in sharing more alleles with populations from Neolithic Iran than Neolithic Anatolia and Europe. (d) Bronze Age Anatolians differ from Mycenaeans in sharing more alleles with Neolithic and Bronze Age populations of the Levant.
Extended Data Figure 4
Extended Data Figure 4. f3-statistics of Mycenaeans as a target with different pairs of reference populations
We show the value of the statistic f3(Ref1, Ref2; Mycenaean) and ±3 standard errors; only the population pairs (Ref1, Ref2) for which the Z-score of the statistic is <−2 are shown. Negative values indicate that the Mycenaean population is admixed from sources related to the two reference populations.
Extended Data Figure 5
Extended Data Figure 5. Correspondence of qpAdm estimates with PCA
As a way to validate qpAdm models of admixture for Myceneans from three ancestral populations (Anatolia_N or Minoan_Lasithi), (Armenia_ChL or Armenia_MLBA), (Steppe_EMBA, Steppe_MLBA, Europe_LNBA), representing substratum, ‘eastern’, and ‘northern’ ancestry respectively (Supplementary Information, section 2), we plot the qpAdm-predicted position in the PCA space of Fig. 1 vs. the actual position of the Mycenaean population.
Extended Data Figure 6
Extended Data Figure 6. Comparison of Mycenaeans and simulated admixed populations
We simulate admixed individuals with known ancestry from three ancestral populations (Anatolia_N or Minoan_Lasithi), (Armenia_ChL or Armenia_MLBA), (Steppe_EMBA, Steppe_MLBA, Europe_LNBA), representing substratum, ‘eastern’, and ‘northern’ ancestry respectively (Methods; Supplementary Information, section 2). The maximum |Z|-score of statistics f4(Mycenaean, Simulated; Outgroup1, Outgroup2) is plotted with circles of varying size (proportional to log|Z|) for each assignment of ancestry proportions. The best estimate (red) corresponds to the proportions that minimize |Z|, and they are compared against the qpAdm estimate for the same ancestral sources (blue).
Extended Data Figure 7
Extended Data Figure 7. FST between Bronze Age and present-day West Eurasian populations
(a) The population of Early Bronze Age Armenia shows an affinity to present-day populations from Armenia, Anatolia, the Caucasus, and Iran, as does (b) Middle/Late Bronze Age Armenia,. (c) The Bronze Age Levant has an affinity to Levantine and Arabian populations. (d) Late Neolithic/Bronze Age Europeans,,, most resemble present-day northern/central Europeans, as do (e) Early/Middle Bronze Age steppe populations,,, who also resemble populations of the northeast Caucasus, while (f) Middle/Late Bronze Age steppe populations resemble central/northern Europeans,. Jewish populations are plotted with a square to distinguish them from non-Jewish populations from the same geographical area. The plots for the newly reported populations of Mycenaeans, Minoans, and Bronze Age Anatolians are shown in Fig. 2.
Extended Data Figure 8
Extended Data Figure 8. Symmetry testing of Mycenaeans with Modern Greek populations
The statistic f4(Mycenaean, Modern Greek; Test, Chimp) is shown with ±3 standard errors. Modern Greeks share fewer alleles with Levantine/Anatolian/European Neolithic populations and with Minoans than Mycenaeans do, suggesting a dilution of early Neolithic ancestry since the Bronze Age. Human Origins genotype data: (a) Greeks from the Coriell repository, (b) Greeks from Thessaloniki, (c) Cypriots. Whole genome data: (d) Cretans. Illumina genotype data: (e) Greeks from Thessaly, (f) Greeks from Central Greece, (g) Greeks from the study by Hellenthal et al.
Figure 1
Figure 1. Samples and principal components analysis
(a) Geographical locations of newly reported ancient data. Lines point to sampling locations; jitter is added to show the number of sampled individual per location. (b) 334 ancient individuals projected onto the first two principal components computed on a sample of 1,029 present-day West Eurasians,,,, including 30 Modern Greek samples from Greece and Cyprus.
Figure 2
Figure 2. Genetic differentiation of Bronze Age populations to present-day populations
We plot the FST inbreeding coefficient (Methods) between newly reported populations and present-day West Eurasian populations which shows a pattern of genetic affinity between Bronze Age and present-day populations from the corresponding broad geographical regions. (a) Mycenaeans, (b) Minoans from Hagios Charalambos (Lasithi regional unit), (c) Minoans from Moni Odigitria (Heraklion regional unit), (d) southwestern Bronze Age Anatolians. The same pattern also applies to Bronze Age populations from other regions of West Eurasia (Extended Data Fig. 5).

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